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Draconology © VikasRao

Please do not copy, redistribute, reference, trace, reuse, alter or modify for your own use without my prior permission, thank you!

Give it a second to load, it's a big GIF! I wanted to try something a little different for this one, mostly as an experiment to see if I could apply it to all future posts going forward, and is why I couldn't post this for #smaugust and instead had to put it off for #spectember so let me know what you think. If it works out, I can imagine showing everything from colour variations to anatomical differences in a single uncluttered image.
The biggest challenge with this was no doubt trying to avoid using too many bird references during the design process but still making sure they were recognizable as gryphons. So I used turtle and ceratopsian beaks and heads, but soon realized no matter how hard I tried, the only way to keep the gryphon aesthetic was to retain some avian design influences, but I figured that as long as it was restricted to the head, it would be just fine. After all, beaks aren't exactly exclusive to birds!
……………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………

The Gryphonidae is a family of ornithodraconian bird dragons consisting of 12 species that are traditionally referred to as gryphons, griffins and griffons. While they bear a passing resemblance to birds of prey and azhdarchid pterosaurs, this is merely a case of convergent evolution, as they are unrelated taxa and last shared a common ancestor over 300 mya. Within Ornithodraconia, gryphons belong to the superfamily Ornithodraconoidea, alongside the cockatrices (Ornithodraconidae) & 2 extinct groups. The superficial resemblance to birds is a result of their thick, downy coelofibers and a keratinous beak, which is actually formed by the fusion of several rostral osteoderms similar to the beaks of turtles and tortoises rather than birds or pterosaurs.

While cockatrices & gryphons are anatomically very similar, they are relatively easy to distinguish: gryphons are generally larger, have larger heads with raptorial, hooked beaks, lack teeth and, with the exception of 2 species, are obligate carnivores, whereas cockatrices are typically generalist omnivores. The hooked tip is actually a keratin-covered premaxillary tooth. While all known dragon species do possess a premaxillary tooth/egg tooth as embryos and hatchlings, these are lost fairly quickly, but in gryphons, this trait is retained well into adulthood and modified to serve as a weapon. None of the known gryphon species possess teeth or ridges like their Ornithodraconid cousins, but their hooked beaks have more than made up for this. Barring 2 omnivorous species, all other gryphons are obligate carnivores, feeding on just about everything from fish to carrion to medium & large game.

In contrast to the true dragons (Eudraconia) which show a high degree of morphological conservation, gryphonids show a considerable amount of morphological variation. This is likely a result of specialization to fit into distinct niches not occupied by other taxa in their range, that has allowed the surviving species to persist. The smallest gryphons are no bigger than an eagle, while the largest rival some true dragons, although none are as large as the largest true dragons. All gryphons are quite agile on the ground, as with most Ornithodraconians, and some are even arboreal.

Fossil evidence suggests that gryphons were the first macropredatory draconians to evolve in the aftermath of the K-Pg extinction, likely due to their faster rate of reproduction. They split off from their ornithodraconid cousins roughly 60 mya in the Paleocene epoch, and took over the role of apex predators, a niche that had been left vacant since the extinction of the non-avian dinosaurs. They held this position until the end of the Eocene, when they were largely outcompeted by the Eudraconoid dragons, mammalian predators, and the teratorns and thunderbirds (in the Americas). Although gryphons have disappeared from the Americas, they continue to survive on the other continents, with one species even having successfully established itself in Antarctica. While the Gryphonidae are the only extant lineage of macropredatory ornithodraconians, fossil evidence shows that they're hardly unique, with many extinct taxa from the other ornithodraconoidean lineages having evolved very similar traits and presumably lifestyles. Notably, in the Americas where gryphons never truly established, a few species of cockatrice have convergently evolved many gryphon-like traits and occupy similar niches as the smaller gryphon species.

Macropredatory gryphons appear to have been present in Europe - particularly on islands - until as recently as the last glacial maximum around 26,000 years ago. This was likely due to 2 factors: many of the European islands were sufficiently isolated from the mainland to allow these gryphons to persist, and that prior to this period, prey populations appear to have been large enough to sustain mammalian predators, gryphons and dragons. However, as sea levels continued to drop during the Ice Age, mammalian predators were able to colonize several islands due to newly formed land bridges, which, coupled with drastic prey population declines during the last glacial maximum, resulted in intensified competition between the 3 predator classes. The mammalian predators that survived mostly managed to do so thanks to their more generalist habits & faster reproductive rates compared to their draconian competitors, & dragons likely survived due to them being better adapted to hunting larger prey, as well as their higher intelligence and fire breathing traits.

The surviving gryphon species are widespread across much of Eurasia, Africa, Australia & the surrounding islands, but are absent from the Americas. While 2 fossil taxa are known from the Paleogene of North America, the gryphon fossil record in South America is non-existent, suggesting that they probably never managed to colonize the continent at any point in time.

The 12 species of gryphons are classified into 3 distinct subfamilies: the Hesperogryphoninae, Gryphoninae & Sagittariopterinae.

Weighing in at a hefty 100-115 kg and standing slightly over 2m in height, the Atlantean giant alke (Pachygryphus robustus) is the heaviest extant gryphon, and with a wingspan of less than 5 m, the only known flightless species. Endemic to Atlantis, it is the sole surviving member of the subfamily Hesperogryphoninae, a clade of basal gryphons that included the only gryphonids known to have reached North America. The alke's presence in Atlantis predates that of its extinct North American relatives, implying that the islands may have served as a bridge allowing them to disperse west.

In contrast to other gryphons, the alke is hypocarnivorous, i.e, most of its diet comprises of plants rather than meat, similar to many bear species. Indeed, its large size is mostly due to its large and rather complex gut, an adaptation to feeding on fruit, fresh leaves, roots & tubers. Most meat consumed is usually in the form of carrion, eggs, and bones, which it can break open with its powerful beak. Juveniles are more carnivorous than adults, and actively hunt small prey such as arthropods & small vertebrates. Both sexes are similar in size, and unusually among gryphons, this species mates for life. The female lays 4-6 eggs every few years, which are incubated solely by the male for nearly 2 months. The offspring (flaplings) though are looked after by both parents for over 3 years.

The Gryphoninae is the most diverse subfamily, comprised of 7 species in 2 genera. The group is further divided into 2 distinct tribes: Gryphonini & Hemigryphonini, with 5 species in the former and 2 in the latter. Fossils show that this subfamily was hit hard by the Middle Miocene extinction around 14 mya, reducing 5 distinct tribes to 2.

By far the best-known member of this clade is the garuda (Gryphon melanops) or Asiatic gryphon. With a wingspan of 5 – 6 m, it isn’t the largest gryphon, but with a range extending from Western Europe through virtually all of Asia, is easily the most widespread. A gracile gryphon, it inhabits open steppe, grasslands and scrub forests, preying upon small game such as rodents, lagomorphs, small primates and young ungulates, as well as scavenging upon carrion. Because of its dietary preference, it is able to coexist with the many species of true dragons that also inhabit the continent, as these generally hunt larger game. in addition to being among the fastest running of all gryphons, garudas are also quite aerobatic fliers, which helps them evade attacks from these dragons. Although generally non-aggressive, they will not retreat easily, only giving way to the very largest or fiercest of beasts. During the breeding season however, males can become especially aggressive and ill-tempered. Garudas don’t mate for life, but like many true dragons form monogamous pairings with females for the duration of the breeding season, and males go from their regular dark brown to a striking maroon and white colouration during this time and engage in spectacular fights. These fights can be quite intense, and combatants may sometimes sustain fatal injuries. Pumped with testosterone, breeding males may even attack large dragons and anything else that may get too close.

This species first appeared 9 mya in the late Miocene, making it the oldest surviving gryphoninin. Phylogenetic studies actually suggest that the garuda sits at the very base of the Gryphonini phylogenetic tree, and is ancestral to the other 4 species within the tribe. It is thought that following the near extinction of this subfamily, the garuda became widespread through most of Eurasia, even reaching most Southeast Asian islands, and subsequently giving rise to several species.

The Opinicus (Gryphon aquiloides) or European spotted gryphon is very closely related to the garuda, so much so that the 2 species are able to produce fertile hybrids. Indeed, western garuda populations show considerable genetic admixture with opinicuses, and eastern opinicus populations show high degree of admixture with garudas. Genetic evidence indicates that a distinct European population of garuda became separated from the main Asian population during the Pleistocene likely as a result of glacial expansion, and gave rise to a distinct species over the course of the next 1million years. It appears that the main reason the 2 species have remained distinct despite the retreat of glaciers is their slightly differing habitat preference and breeding chronology. Because both species breed in the spring, more northerly populations of opinicus breed later compared than central and southern ones, which breed around the same time as central and West Asian garuda populations. However, opinici in general have a greater preference for woodlands compared to garudas, which prefer open habitats. This results in niche partitioning in regions where the 2 species coexist, so while encounters do occur, matings are much less frequent.

Opinici also spend more time hunting from the ground than from the air, feeding on small prey such as rodents and birds as well as deer fawns and carrion. Although they don't display breeding colours like garudas, opinici do show sexual dimorphism. Both sexes are of similar size, and start out with a spotted pelage that provides camouflage in their forested habitat. As they become juveniles, they lose their spots and turn a uniform brown colour. Females will retain this pelage their entire lives, but males, upon reaching maturity, will revert back to their spotted pelage. During the breeding season, nesting opinici are known to associate to varying extents with the ocellated wyvern (Volanosaurus ocellatus). Because both parents guard the nest at night, the smaller wyvern will oftentimes nest in close proximity so that its own offspring are protected at as it forages. It is unclear if the gryphon benefits from this association, but it's believed that the wyvern keeps the nest free of parasites and pests.

The only extant species of macropredatory gryphon is the southern hippogryph (Gryphon ischyrorhynchos) of Aotearoa. Standing 2m tall with a wingspan just under 7m and weighing 65-70 kg, this is the largest member of the Gryphoninae, and one of the largest extant gryphons. Although capable of flight, it spends most of its time on the ground, only taking to the air to travel large distances. A heavily built animal with a thick, robust beak, the hippogryph hunts giant moas and other large flightless birds of the island, using its powerful wings to launch itself forward and delivering a debilitating strike to its prey's head or neck, killing it rapidly before it has a chance to outrun it. Juvenile hippogryphs, which are more agile than adults, feed on smaller prey such as waterfowl.

Because dragons never reached Aotearoa, the southern hippogryph has remained the apex predator of the island, though it itself appears to be a rather recent arrival, having colonized the island just over 4 million years ago in the early Pliocene. Its only competitor today is the pouakai (Hieraaetus moorei), a giant eagle that arrived much more recently in the mid Pleistocene. The 2 predators do manage to coexist however due to differential habitat preference and niche partitioning: the eagle, a diurnal aerial divebomber, prefers more open habitats & highlands, whereas the hippogryph, a crepuscular terrestrial ambush hunter, prefers much denser woodland, and due to its larger size, generally tackles larger prey than the eagle. Nonetheless, though encounters between the 2 are infrequent, they aren't unheard of. Pouakai can kill hippogryphs by divebombing them, should they manage to catch one unawares, and hippogryphs are more than capable of killing an eagle on the ground, and both predators will oftentimes steal each other's kills.

The 2 surviving hemigryphoninin species represent the last of a basal lineage of gryphonines, which, in terms of general appearance, bear a strong resemblance to their distant eudraconian (true dragon) cousins.

The lesser keythong (Hemigryphon primaevus), also called the lesser horned gryphon, is the smallest gryphon species, with a wingspan of roughly 1.7-2 m. Endemic to Madagascar, this species is notable for its backward-pointing cranial horns, spiny quills on its neck, back and tail, and large osteoderms on its head that give it an armored appearance. They are also known to be particularly foul-tempered animals, and will rarely back down from a confrontation even against foes several times their own size, and have been known to fight viciously if cornered or if their offspring are threatened. Their distinctive white heads, black bodies and barred quills appear to be part of an aposematic display; a warning to other animals to keep their distance.

The reason for these seemingly excessive defences and aggression is the keythong's habitat and lifestyle: it inhabits Madagascar's forests, where it forages on the ground for invertebrates and small vertebrates, and while its size allows it to survive on the limited resources of its island habitat, it also makes it vulnerable to ambush by fossas, crocodiles, eagles, and dragons, as well as harassment by lemurs. Although capable of flight, it isn't always an option in a dense habitat where most threats are either arboreal or similarly flight-capable. Indeed, a close relative, the greater keythong (Hemigryphon trigonocephalus) of eastern Africa which inhabits open savannah and sparser woodlands, has smaller quills but is larger in size.

These gryphons are also unusual in that they seem to be a K-selected species, with females laying 2 eggs once every 2 years that are looked after by both parents. Adults generally form lifelong monogamous pairs and care for their offspring for over a year, because of which, juvenile mortality is rather low.

The 4 species of the Sagittariopterinae are commonly called axexes, and appear to have descended from a lineage of gryphons that were at least partially semi-aquatic, as all extant species are tied to water bodies to varying degrees.

With a 3-4 m wingspan and standing just under 1.3 m in height, the Southern axex (Axex australis) is one of the smaller members of this subfamily, though is still comparable in size to a very large albatross. As its name suggests, it inhabits much of the Southern Hemisphere, from the southern tips of Africa and South America to Australia and even Antarctica, making it the only draconian, and indeed the only non-avian reptile, that has a sizeable population in the Antarctic. As with its larger cousin the Northern axex (Axex borealis), this species has a seabird-like lifestyle, spending most of its time dynamically soaring over the ocean, sometimes for days at a time, as it hunts on the wing. Oddly enough, it occupies a similar ecological niche as a sea-going vulture, feeding primarily on the carcasses of marine mammals and seabirds, and only hunting fish and squid on occasion. Its large size & aggression allows it to drive away scavenging seabirds such as giant petrels and gulls, but at the same time makes carcasses more accessible to these smaller scavengers; its powerful hooked beak is better at tearing open the thick hides of seals and whales, allowing the birds to access more meat. For this reason, many smaller scavengers such as gulls will intentionally alert a passing axex to the presence of a carcass.

Southern axexes mate in late winter, and females lay around 6-10 eggs shortly before the summer. Breeding adults will form monogamous pairs for the duration of the breeding season, and both parents incubate the eggs until they hatch. The hatching of the eggs usually coincides with the beginning of summer, and the arrival of several pinniped and seabird species. As with most other gryphons, axex flaplings can fly within a few days of hatching, and a calorie-rich diet of blubbery prey & carcasses allows them to grow at a rapid rate, reaching adult size in as little as 3-4 months, though not many survive to this age, as they are oftentimes killed by everything from predatory seals, orcas, groups of petrels & skuas and other axexes.

The painted axex (Sagittariopteryx regalis) is a medium-sized gryphon native to northeastern and much of Sub-Saharan Africa. With a wingspan of 5-5.5m, it is similar in size to the garuda, and occupies a similar ecological niche as a small-game hunter. With a maximum weight of 35 kg however, the axex is a more heavily-built animal, and therefore isn't quite as aerobatic as its Asian counterpart. At the same time, although it's a habitat generalist, it shows a strong preference for habitats with large water bodies and a corresponding greater reliance on fish, amphibians & reptiles.

While not particularly agile in the air, the painted axex is, even by gryphon standards, quite agile on the ground, to the point that it will chase birds and rodents and even attempt to evade predators by running away, only taking to the air as a last resort. In fact, compared to most gryphons, the axex will opt for fight over flight in almost half of all encounters with predators, only fleeing when faced with much larger predators. In this regard, the painted axex is quite dragon-like in terms of behaviour. While not a social species, they will generally forage in loose aggregations or groups, which will cooperate to mob smaller predators such as jackals, cheetahs, lone hyenas and even leopards and smaller dragons, and will regularly hunt other reptilian predators such as large pythons (though a lone individual can easily fall prey to any one of these). The few predators that axexes are known to flee from are lions, large dragons and packs of African wild dogs and spotted hyenas.

Standing over 2 m tall ith a 7-8 m wingspan and weighing upto 75-85 kg, the Lemurian giant axex (Sagittariopteryx giganteus) is the largest extant gryphon species. Larger than both the hippogryph and the Atlantean alke (though not as heavy as the latter), this species is endemic to Lemuria, and appears to be the sister species of the painted axex, the 2 having diverged from a common ancestor over 6 mya. The giant axex is unique among gryphons in being a wader, with a feeding ecology similar to large wading birds such as storks and herons. However, it doesn't rely as much on fish as the birds, and instead, feeds primarily on molluscs such as freshwater snails, including several species of giant snails that inhabit Lemuria's river systems. Smaller snails are generally devoured whole, while larger species with tougher shells are cracked open either by hitting them with rocks or by dropping them from the air. The tip of its tongue is covered in large keratinous papillae, which allow it to grip and scrape the soft parts of its prey. Although snails make up 65% of its diet, it will also regularly hunt fish, amphibians even juvenile crocodylians, grabbing them with its large, broad bill and either swallowing then whole or killing them by beating them on the ground. This species is the only other omnivorous gryphon besides the Atlantean alke, and will occasionally feed on fruit, podocarp cones and tubers. In particular, it shows a strong preference for the fruit of the Pagoda tree, a species of Myristica endemic to Lemuria. This particular tree only bears fruit once every 5-6 years, an event that offers bountiful pickings for the region's herbivores. The fruit appears to play an important role in the gryphon's ecology, as fruiting years generally see higher offspring survival than non-fruiting years, likely due to the fruit's high nutritional value and the relative ease of accessibility. In turn, the giant axex appears to be one of the plant's most important dispersal agents. During the dry season, these gryphons forage in wetlands and mangroves, but during the wet season will forage in floodplains, river basins and highlands, traversing the entire microcontinent as a result and dispersing the seeds as they go.

Lemurian giant axexes are sexually dimorphic, with males generally averaging slightly larger than females. As with most gryphons, they don't mate for life, but generally form monogamous pairs for the duration of the breeding season, although extra pair matings are not unheard of. Oddly enough, only the male looks after the flaplings, defending them against predators for 3-4 months. Both sexes start out with dark grey to near black pelage, but as males mature (around 8-10 years of age), the coelofibers of their necks turn white and the rest develop maroon-burgundy highlights.

Draconology: The Ornithodraconia

Draconology: The Draconimorpha

Edit: Minor redesign of the alke to better reflect my original idea of it as an owlbear-type gryphon.

Description

Draconology © VikasRao

Please do not copy, redistribute, reference, trace, reuse, alter or modify for your own use without my prior permission, thank you!

Give it a second to load, it's a big GIF! I wanted to try something a little different for this one, mostly as an experiment to see if I could apply it to all future posts going forward, and is why I couldn't post this for #smaugust and instead had to put it off for #spectember so let me know what you think. If it works out, I can imagine showing everything from colour variations to anatomical differences in a single uncluttered image.
The biggest challenge with this was no doubt trying to avoid using too many bird references during the design process but still making sure they were recognizable as gryphons. So I used turtle and ceratopsian beaks and heads, but soon realized no matter how hard I tried, the only way to keep the gryphon aesthetic was to retain some avian design influences, but I figured that as long as it was restricted to the head, it would be just fine. After all, beaks aren't exactly exclusive to birds!
……………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………………

The Gryphonidae is a family of ornithodraconian bird dragons consisting of 12 species that are traditionally referred to as gryphons, griffins and griffons. While they bear a passing resemblance to birds of prey and azhdarchid pterosaurs, this is merely a case of convergent evolution, as they are unrelated taxa and last shared a common ancestor over 300 mya. Within Ornithodraconia, gryphons belong to the superfamily Ornithodraconoidea, alongside the cockatrices (Ornithodraconidae) & 2 extinct groups. The superficial resemblance to birds is a result of their thick, downy coelofibers and a keratinous beak, which is actually formed by the fusion of several rostral osteoderms similar to the beaks of turtles and tortoises rather than birds or pterosaurs.

While cockatrices & gryphons are anatomically very similar, they are relatively easy to distinguish: gryphons are generally larger, have larger heads with raptorial, hooked beaks, lack teeth and, with the exception of 2 species, are obligate carnivores, whereas cockatrices are typically generalist omnivores. The hooked tip is actually a keratin-covered premaxillary tooth. While all known dragon species do possess a premaxillary tooth/egg tooth as embryos and hatchlings, these are lost fairly quickly, but in gryphons, this trait is retained well into adulthood and modified to serve as a weapon. None of the known gryphon species possess teeth or ridges like their Ornithodraconid cousins, but their hooked beaks have more than made up for this. Barring 2 omnivorous species, all other gryphons are obligate carnivores, feeding on just about everything from fish to carrion to medium & large game.

In contrast to the true dragons (Eudraconia) which show a high degree of morphological conservation, gryphonids show a considerable amount of morphological variation. This is likely a result of specialization to fit into distinct niches not occupied by other taxa in their range, that has allowed the surviving species to persist. The smallest gryphons are no bigger than an eagle, while the largest rival some true dragons, although none are as large as the largest true dragons. All gryphons are quite agile on the ground, as with most Ornithodraconians, and some are even arboreal.

Fossil evidence suggests that gryphons were the first macropredatory draconians to evolve in the aftermath of the K-Pg extinction, likely due to their faster rate of reproduction. They split off from their ornithodraconid cousins roughly 60 mya in the Paleocene epoch, and took over the role of apex predators, a niche that had been left vacant since the extinction of the non-avian dinosaurs. They held this position until the end of the Eocene, when they were largely outcompeted by the Eudraconoid dragons, mammalian predators, and the teratorns and thunderbirds (in the Americas). Although gryphons have disappeared from the Americas, they continue to survive on the other continents, with one species even having successfully established itself in Antarctica. While the Gryphonidae are the only extant lineage of macropredatory ornithodraconians, fossil evidence shows that they're hardly unique, with many extinct taxa from the other ornithodraconoidean lineages having evolved very similar traits and presumably lifestyles. Notably, in the Americas where gryphons never truly established, a few species of cockatrice have convergently evolved many gryphon-like traits and occupy similar niches as the smaller gryphon species.

Macropredatory gryphons appear to have been present in Europe - particularly on islands - until as recently as the last glacial maximum around 26,000 years ago. This was likely due to 2 factors: many of the European islands were sufficiently isolated from the mainland to allow these gryphons to persist, and that prior to this period, prey populations appear to have been large enough to sustain mammalian predators, gryphons and dragons. However, as sea levels continued to drop during the Ice Age, mammalian predators were able to colonize several islands due to newly formed land bridges, which, coupled with drastic prey population declines during the last glacial maximum, resulted in intensified competition between the 3 predator classes. The mammalian predators that survived mostly managed to do so thanks to their more generalist habits & faster reproductive rates compared to their draconian competitors, & dragons likely survived due to them being better adapted to hunting larger prey, as well as their higher intelligence and fire breathing traits.

The surviving gryphon species are widespread across much of Eurasia, Africa, Australia & the surrounding islands, but are absent from the Americas. While 2 fossil taxa are known from the Paleogene of North America, the gryphon fossil record in South America is non-existent, suggesting that they probably never managed to colonize the continent at any point in time.

The 12 species of gryphons are classified into 3 distinct subfamilies: the Hesperogryphoninae, Gryphoninae & Sagittariopterinae.

Weighing in at a hefty 100-115 kg and standing slightly over 2m in height, the Atlantean giant alke (Pachygryphus robustus) is the heaviest extant gryphon, and with a wingspan of less than 5 m, the only known flightless species. Endemic to Atlantis, it is the sole surviving member of the subfamily Hesperogryphoninae, a clade of basal gryphons that included the only gryphonids known to have reached North America. The alke's presence in Atlantis predates that of its extinct North American relatives, implying that the islands may have served as a bridge allowing them to disperse west.

In contrast to other gryphons, the alke is hypocarnivorous, i.e, most of its diet comprises of plants rather than meat, similar to many bear species. Indeed, its large size is mostly due to its large and rather complex gut, an adaptation to feeding on fruit, fresh leaves, roots & tubers. Most meat consumed is usually in the form of carrion, eggs, and bones, which it can break open with its powerful beak. Juveniles are more carnivorous than adults, and actively hunt small prey such as arthropods & small vertebrates. Both sexes are similar in size, and unusually among gryphons, this species mates for life. The female lays 4-6 eggs every few years, which are incubated solely by the male for nearly 2 months. The offspring (flaplings) though are looked after by both parents for over 3 years.

The Gryphoninae is the most diverse subfamily, comprised of 7 species in 2 genera. The group is further divided into 2 distinct tribes: Gryphonini & Hemigryphonini, with 5 species in the former and 2 in the latter. Fossils show that this subfamily was hit hard by the Middle Miocene extinction around 14 mya, reducing 5 distinct tribes to 2.

By far the best-known member of this clade is the garuda (Gryphon melanops) or Asiatic gryphon. With a wingspan of 5 – 6 m, it isn’t the largest gryphon, but with a range extending from Western Europe through virtually all of Asia, is easily the most widespread. A gracile gryphon, it inhabits open steppe, grasslands and scrub forests, preying upon small game such as rodents, lagomorphs, small primates and young ungulates, as well as scavenging upon carrion. Because of its dietary preference, it is able to coexist with the many species of true dragons that also inhabit the continent, as these generally hunt larger game. in addition to being among the fastest running of all gryphons, garudas are also quite aerobatic fliers, which helps them evade attacks from these dragons. Although generally non-aggressive, they will not retreat easily, only giving way to the very largest or fiercest of beasts. During the breeding season however, males can become especially aggressive and ill-tempered. Garudas don’t mate for life, but like many true dragons form monogamous pairings with females for the duration of the breeding season, and males go from their regular dark brown to a striking maroon and white colouration during this time and engage in spectacular fights. These fights can be quite intense, and combatants may sometimes sustain fatal injuries. Pumped with testosterone, breeding males may even attack large dragons and anything else that may get too close.

This species first appeared 9 mya in the late Miocene, making it the oldest surviving gryphoninin. Phylogenetic studies actually suggest that the garuda sits at the very base of the Gryphonini phylogenetic tree, and is ancestral to the other 4 species within the tribe. It is thought that following the near extinction of this subfamily, the garuda became widespread through most of Eurasia, even reaching most Southeast Asian islands, and subsequently giving rise to several species.

The Opinicus (Gryphon aquiloides) or European spotted gryphon is very closely related to the garuda, so much so that the 2 species are able to produce fertile hybrids. Indeed, western garuda populations show considerable genetic admixture with opinicuses, and eastern opinicus populations show high degree of admixture with garudas. Genetic evidence indicates that a distinct European population of garuda became separated from the main Asian population during the Pleistocene likely as a result of glacial expansion, and gave rise to a distinct species over the course of the next 1million years. It appears that the main reason the 2 species have remained distinct despite the retreat of glaciers is their slightly differing habitat preference and breeding chronology. Because both species breed in the spring, more northerly populations of opinicus breed later compared than central and southern ones, which breed around the same time as central and West Asian garuda populations. However, opinici in general have a greater preference for woodlands compared to garudas, which prefer open habitats. This results in niche partitioning in regions where the 2 species coexist, so while encounters do occur, matings are much less frequent.

Opinici also spend more time hunting from the ground than from the air, feeding on small prey such as rodents and birds as well as deer fawns and carrion. Although they don't display breeding colours like garudas, opinici do show sexual dimorphism. Both sexes are of similar size, and start out with a spotted pelage that provides camouflage in their forested habitat. As they become juveniles, they lose their spots and turn a uniform brown colour. Females will retain this pelage their entire lives, but males, upon reaching maturity, will revert back to their spotted pelage. During the breeding season, nesting opinici are known to associate to varying extents with the ocellated wyvern (Volanosaurus ocellatus). Because both parents guard the nest at night, the smaller wyvern will oftentimes nest in close proximity so that its own offspring are protected at as it forages. It is unclear if the gryphon benefits from this association, but it's believed that the wyvern keeps the nest free of parasites and pests.

The only extant species of macropredatory gryphon is the southern hippogryph (Gryphon ischyrorhynchos) of Aotearoa. Standing 2m tall with a wingspan just under 7m and weighing 65-70 kg, this is the largest member of the Gryphoninae, and one of the largest extant gryphons. Although capable of flight, it spends most of its time on the ground, only taking to the air to travel large distances. A heavily built animal with a thick, robust beak, the hippogryph hunts giant moas and other large flightless birds of the island, using its powerful wings to launch itself forward and delivering a debilitating strike to its prey's head or neck, killing it rapidly before it has a chance to outrun it. Juvenile hippogryphs, which are more agile than adults, feed on smaller prey such as waterfowl.

Because dragons never reached Aotearoa, the southern hippogryph has remained the apex predator of the island, though it itself appears to be a rather recent arrival, having colonized the island just over 4 million years ago in the early Pliocene. Its only competitor today is the pouakai (Hieraaetus moorei), a giant eagle that arrived much more recently in the mid Pleistocene. The 2 predators do manage to coexist however due to differential habitat preference and niche partitioning: the eagle, a diurnal aerial divebomber, prefers more open habitats & highlands, whereas the hippogryph, a crepuscular terrestrial ambush hunter, prefers much denser woodland, and due to its larger size, generally tackles larger prey than the eagle. Nonetheless, though encounters between the 2 are infrequent, they aren't unheard of. Pouakai can kill hippogryphs by divebombing them, should they manage to catch one unawares, and hippogryphs are more than capable of killing an eagle on the ground, and both predators will oftentimes steal each other's kills.

The 2 surviving hemigryphoninin species represent the last of a basal lineage of gryphonines, which, in terms of general appearance, bear a strong resemblance to their distant eudraconian (true dragon) cousins.

The lesser keythong (Hemigryphon primaevus), also called the lesser horned gryphon, is the smallest gryphon species, with a wingspan of roughly 1.7-2 m. Endemic to Madagascar, this species is notable for its backward-pointing cranial horns, spiny quills on its neck, back and tail, and large osteoderms on its head that give it an armored appearance. They are also known to be particularly foul-tempered animals, and will rarely back down from a confrontation even against foes several times their own size, and have been known to fight viciously if cornered or if their offspring are threatened. Their distinctive white heads, black bodies and barred quills appear to be part of an aposematic display; a warning to other animals to keep their distance.

The reason for these seemingly excessive defences and aggression is the keythong's habitat and lifestyle: it inhabits Madagascar's forests, where it forages on the ground for invertebrates and small vertebrates, and while its size allows it to survive on the limited resources of its island habitat, it also makes it vulnerable to ambush by fossas, crocodiles, eagles, and dragons, as well as harassment by lemurs. Although capable of flight, it isn't always an option in a dense habitat where most threats are either arboreal or similarly flight-capable. Indeed, a close relative, the greater keythong (Hemigryphon trigonocephalus) of eastern Africa which inhabits open savannah and sparser woodlands, has smaller quills but is larger in size.

These gryphons are also unusual in that they seem to be a K-selected species, with females laying 2 eggs once every 2 years that are looked after by both parents. Adults generally form lifelong monogamous pairs and care for their offspring for over a year, because of which, juvenile mortality is rather low.

The 4 species of the Sagittariopterinae are commonly called axexes, and appear to have descended from a lineage of gryphons that were at least partially semi-aquatic, as all extant species are tied to water bodies to varying degrees.

With a 3-4 m wingspan and standing just under 1.3 m in height, the Southern axex (Axex australis) is one of the smaller members of this subfamily, though is still comparable in size to a very large albatross. As its name suggests, it inhabits much of the Southern Hemisphere, from the southern tips of Africa and South America to Australia and even Antarctica, making it the only draconian, and indeed the only non-avian reptile, that has a sizeable population in the Antarctic. As with its larger cousin the Northern axex (Axex borealis), this species has a seabird-like lifestyle, spending most of its time dynamically soaring over the ocean, sometimes for days at a time, as it hunts on the wing. Oddly enough, it occupies a similar ecological niche as a sea-going vulture, feeding primarily on the carcasses of marine mammals and seabirds, and only hunting fish and squid on occasion. Its large size & aggression allows it to drive away scavenging seabirds such as giant petrels and gulls, but at the same time makes carcasses more accessible to these smaller scavengers; its powerful hooked beak is better at tearing open the thick hides of seals and whales, allowing the birds to access more meat. For this reason, many smaller scavengers such as gulls will intentionally alert a passing axex to the presence of a carcass.

Southern axexes mate in late winter, and females lay around 6-10 eggs shortly before the summer. Breeding adults will form monogamous pairs for the duration of the breeding season, and both parents incubate the eggs until they hatch. The hatching of the eggs usually coincides with the beginning of summer, and the arrival of several pinniped and seabird species. As with most other gryphons, axex flaplings can fly within a few days of hatching, and a calorie-rich diet of blubbery prey & carcasses allows them to grow at a rapid rate, reaching adult size in as little as 3-4 months, though not many survive to this age, as they are oftentimes killed by everything from predatory seals, orcas, groups of petrels & skuas and other axexes.

The painted axex (Sagittariopteryx regalis) is a medium-sized gryphon native to northeastern and much of Sub-Saharan Africa. With a wingspan of 5-5.5m, it is similar in size to the garuda, and occupies a similar ecological niche as a small-game hunter. With a maximum weight of 35 kg however, the axex is a more heavily-built animal, and therefore isn't quite as aerobatic as its Asian counterpart. At the same time, although it's a habitat generalist, it shows a strong preference for habitats with large water bodies and a corresponding greater reliance on fish, amphibians & reptiles.

While not particularly agile in the air, the painted axex is, even by gryphon standards, quite agile on the ground, to the point that it will chase birds and rodents and even attempt to evade predators by running away, only taking to the air as a last resort. In fact, compared to most gryphons, the axex will opt for fight over flight in almost half of all encounters with predators, only fleeing when faced with much larger predators. In this regard, the painted axex is quite dragon-like in terms of behaviour. While not a social species, they will generally forage in loose aggregations or groups, which will cooperate to mob smaller predators such as jackals, cheetahs, lone hyenas and even leopards and smaller dragons, and will regularly hunt other reptilian predators such as large pythons (though a lone individual can easily fall prey to any one of these). The few predators that axexes are known to flee from are lions, large dragons and packs of African wild dogs and spotted hyenas.

Standing over 2 m tall ith a 7-8 m wingspan and weighing upto 75-85 kg, the Lemurian giant axex (Sagittariopteryx giganteus) is the largest extant gryphon species. Larger than both the hippogryph and the Atlantean alke (though not as heavy as the latter), this species is endemic to Lemuria, and appears to be the sister species of the painted axex, the 2 having diverged from a common ancestor over 6 mya. The giant axex is unique among gryphons in being a wader, with a feeding ecology similar to large wading birds such as storks and herons. However, it doesn't rely as much on fish as the birds, and instead, feeds primarily on molluscs such as freshwater snails, including several species of giant snails that inhabit Lemuria's river systems. Smaller snails are generally devoured whole, while larger species with tougher shells are cracked open either by hitting them with rocks or by dropping them from the air. The tip of its tongue is covered in large keratinous papillae, which allow it to grip and scrape the soft parts of its prey. Although snails make up 65% of its diet, it will also regularly hunt fish, amphibians even juvenile crocodylians, grabbing them with its large, broad bill and either swallowing then whole or killing them by beating them on the ground. This species is the only other omnivorous gryphon besides the Atlantean alke, and will occasionally feed on fruit, podocarp cones and tubers. In particular, it shows a strong preference for the fruit of the Pagoda tree, a species of Myristica endemic to Lemuria. This particular tree only bears fruit once every 5-6 years, an event that offers bountiful pickings for the region's herbivores. The fruit appears to play an important role in the gryphon's ecology, as fruiting years generally see higher offspring survival than non-fruiting years, likely due to the fruit's high nutritional value and the relative ease of accessibility. In turn, the giant axex appears to be one of the plant's most important dispersal agents. During the dry season, these gryphons forage in wetlands and mangroves, but during the wet season will forage in floodplains, river basins and highlands, traversing the entire microcontinent as a result and dispersing the seeds as they go.

Lemurian giant axexes are sexually dimorphic, with males generally averaging slightly larger than females. As with most gryphons, they don't mate for life, but generally form monogamous pairs for the duration of the breeding season, although extra pair matings are not unheard of. Oddly enough, only the male looks after the flaplings, defending them against predators for 3-4 months. Both sexes start out with dark grey to near black pelage, but as males mature (around 8-10 years of age), the coelofibers of their necks turn white and the rest develop maroon-burgundy highlights.

Draconology: The Ornithodraconia

Draconology: The Draconimorpha

Edit: Minor redesign of the alke to better reflect my original idea of it as an owlbear-type gryphon.